Journal Journal

of Applied
Appl Journal of Applied Horticulture, 22(3): 255-264, 2020 Horticulture
DOI: https://doi.org/10.37855/jah.2020.v22i03.45 ISSN: 0972-1045

Breeding for fusarium wilt resistance and some economic

characters in cucumber

Amani Hafez Abdallah Mahmoud Gharib*, Amr Ahmed El Sayed, Mohamed Adel El Tahawey
and Eman Yehia Khafagi
Horticultural Research Institute and Plant Pathology Research Institute, Agriculture Research Center (ARC), Egypt.
*E-mail: [email protected]

Abstract
This study was conducted during 2017 and 2018 at Kaha Vegetable Research Farm, Horticulture Research Institute, Qalyubia Governorate
to develop promising hybrids of cucumber (Cucumis sativus L.) for yield and fruit quality characters under fusarium wilt (Fusarium
oxysporum) infection stress in Egypt using diallel mating design. The pathogenicity and host range experiment revealed that Fusarium
oxysporum isolate No.3 was the most virulent one to the susceptible cucumber cultivar “Beta alpha” and cucumber was the only infected
host. The prevalence of the non-additive variance suggested heterosis breeding approach is effective way for improvement of these
traits. Most of the traits exhibited significant hybrid vigor for some of crosses based on the better-parent. The analysis of variance
showed that all the studied traits were highly significant indicating that both of the parents and hybrids had high variability. Significant
general and specific combining ability variances were obtained in all studied traits implying that both additive and non-additive gene
effects control genetic expression of these traits. The study showed that lines P2, P4 and P5 had significant positive GCA effects for
total yield trait under fusarium wilt stress. Thus, these parents could be successfully used in future breeding programs. Among all the
crosses, P1×P3, P1×P4, P1×P5, P2×P4 and P2×P5 exhibited significant SCA effects for both early and total yield characters under fusarium
wilt stress. So, these hybrids can be used in future breeding program. The genotypes Kaha1×Dokky2 and Kaha1×Kaha2 followed by
Kaha2×Dokky3 as well as Kaha1×Dokky3 were the most resistant genotypes decreasing disease severity correlated with increase in
total phenol contents and activities of polyphenol oxidase, peroxidase and catalase as compared to susceptible genotypes, i.e., Dokky3,
Kaha1×Dokky1 and Dokky1. The scavenging activity was higher in susceptible genotypes as compared with resistant genotypes.
Furthermore, there was a high correlation between the total phenol content and the scavenging activity. Results also revealed a noticeable
significant correlation between disease severity, total yield and early yield traits. Cluster analysis classified the fifteen genotypes into
five clusters with different number of genotypes. Further improvement of fruit yield could be possible through the hybridization and
selection in transgressive segregation.
Key words: Cucumber, combining ability, heterosis, fusarium wilt, resistance, antioxidant and oxidative activity, DPPH

Introduction diallel analysis had been used often by many cucumber breeders
(Glover et al., 2005; Singh and Asati, 2011; Mule et al., 2012
Cucumber (Cucumis sativus L.) belongs to the family and Ene et al., 2019). Heterosis is an important tool in providing
Cucurbitaceae, which comprises of 117 genera and 825 species breeders to increase yield and its components (Hemant and
with chromosome number 2n=14. It’s one of the most important Tiwari, 2018 and Preethi et al., 2019). Hemant and Tiwari (2018)
and popular cucurbitaceous vegetable crops grown extensively reported that the hybrids with significant heterosis for early
throughout the tropical and subtropical region of the world. yield and the fruit yield per plant is positively associated with
The crop is characterized by high degree of cross-pollination, vegetative characters particularly number of branches per plant
wide range of genetic variability in vegetative and fruit traits. and vine length. Estimates of both general and specific combining
Developing a new cucumber variety is a difficult process and ability are useful in determining the breeding value of cucumber
limited improved varieties have been developed through efficient lines by suggesting their appropriate use in a breeding program.
breeding program targeting high fruit yield and better quality. The The GCA is a measure of the additive gene action, while the SCA
success of any breeding program is determined by collecting is assumed to be dominance gene action, if there is no epistasis.
favorable gene combination in the form of high yield inbreds and Several studies suggested that cucumber can be improved through
heterosis in their crosses (Mule et al., 2012). estimating both general and specific combining abilities (Mule et
al., 2012; Malav et al., 2018 and Prashant et al., 2018).
Combining ability and heterosis are useful tools to a breeder
in any breeding programme. They provide the desired genetic Fusarium wilt of cucumber caused by Fusarium oxysporum f. sp.
description concerning the improvement of crop variety or Cucumerinum (Owen, 1956 and Bedlan, 1986) is a destructive
heterotic exploitation for commercial gains. To be adequately disease accountable for large yield losses. The diagnostic
informed on the heterosis and combining ability of parents in symptoms of Fusarium wilt is a brown discoloration of the
crosses to produce desirable segregating population for selection, vascular system (xylem) when cutting the stems, foliage turn
Journal of Applied Horticulture (www.horticultureresearch.net)
256 Breeding for fusarium wilt resistance and some economic characters in cucumber

yellow, wilting and finally leading to plant death (Owen, 1956; randomized complete design with 3 replications. Each plot was
Gordon and Martyn, 1997; Izzati et al., 2018). The pathogen which 10 plants cultivated by direct seeds in two rows with 0.3×0.3 m
is long-lived in soils, has resulted to use host resistance as the most space between plant to plant. Plants were randomly chosen from
efficient management to avert economic yield loss caused by the each experimental plot at 60 days after sowing to calculate the
disease. Reactive oxygen species (ROS) are chemically reactive disease severity of fusarium wilt.
molecules, form as a natural byproduct of the normal metabolism
Disease assessment: Wilt disease severity (DS) was determined
of oxygen and can increase dramatically and accumulates as a
in both experiments, pot and field according to the following
result of oxidative stress under biotic and abiotic stress conditions
visual scale (0-5) and description as suggested by Grattidge
(Apel and Hirt, 2004; Heller and Tudzynski, 2011). Cells are
and O’Brien (1982). Plants were uprooted and stem and root
able to defend-itself towards damage resulting from ROS, due
were longitudinally dissected for examination of internal tissue
to the action of catalases and peroxidases by scavenging free
discoloration. The wilt disease severity (%) was determined and
radicals (Torres et al., 2006). Also, activity of polyphenol oxidase
calculated using the following formula of Song et al. (2004):
(PPO) increased in response to infection by pathogen (Khatun
(n x v)
et al., 2009; Madadkhah et al., 2012). Polyphenol antioxidants Wilt Disease Severity % = � � × 100
play a function in inhibiting reactive oxygen species (ROS) 5N
Where, n = number of infected plants, v= numerical values of symptoms
harm by scavenging free radicals (Torres et al., 2006), change category, N= total number of plants, 5= maximum numerical value of
cell wall structure, and accumulating antimicrobial secondary symptom catagories. 0: No discoloration, 1: from 1 to 10 % discoloration,
2: from 11 to 30 % discoloration, 3: from 31 to 50 % discoloration, 4:
metabolites which are essential in systemic acquired resistance. from 51 to 75 % discoloration and 5: from 76 % to 100 discoloration.
Plant resistance to biotic and abiotic stresses is frequently
All agronomic cultural practices were applied as recommended
controlled by the metabolic rate of phenolic compounds which
for ordinary cucumber fields. At harvest time, all individual plants
are recognized as hydrophilic antioxidants and play a main part
of five parents and their derived 10 F1 generation were harvested
of the plants defense (Tripathi et al., 2019). DPPH (2,2-diphenyl-
in order to estimate horticultural traits, viz., plant length (cm),
1-picrylhydrazyl) is used to estimate the total phenolic content
and antioxidant activities. Important association between the number of branches per plant, internode length, number of nodes,
compounds and activites showed that phenolic compounds are the fruit length (cm), fruit diameter (cm), fruit weight (g), early yield,
major contributor to the antioxidant properties of these cucumber total yield.
genotypes (Patel and Patel, 2011; Ahmed and Ali, 2013). Sampling, isolation, purification and morphological
identification of the causal pathogen under pot experiment:
The present study aimed to identify a correlation between
Cucumber wilted plants samples were collected from Kaha
differences in the degree of resistance among 15 genotypes of
Research Farm, Horticulture Research Institute, Qalyubia
cucumber against Fusarium wilt disease. Also, bio-chemical
Governorate, Egypt. Isolation from lower stem portions of the
changes in different genotypes were measured and evaluated. In
collected cucumber wilted plants which exhibited different
addition, it aimed to identify breeding lines/varieties having good
degrees of vascular discoloration were rinsed up with tap water
combining ability effects and best cross combinations for developing
for 3 min, and cut into small pieces (1 cm long). The pieces were
promising hybrids with yield and fruit quality characters under
surface sterilized by soaking in 70 % ethanol for 30 s, and placed
fusarium disease stress using diallel mating design and estimating
on filter paper for 2 min to dry out. Four sterilized pieces were
heterosis and phenotypic correlation. transferred onto 9 cm diameter Petri dishes containing 10 mL of
potato dextrose agar medium (PDA) (Difco, 1985). The plates
Materials and methods were incubated at 25±2 °C for 7 days. Pure cultures were obtained
The study was conducted at research facilities of Horticultural by single spore technique. The pure cultures of the growing fungi
Research Institute and Plant Pathology Research Institute, Agric. were examined microscopically and identified (Both, 1971;
Research Center (ARC), Egypt during the period from 2017 to Nelson et al., 1983; Leslie and Summerell, 2006). The isolates
2018 at Kaha Vegetable Research Farm, Qalyubia Governorate, were kept on slants at 4 °C in the refrigerator for further work.
ARC, Egypt. The genetic materials used in this investigation Pathogenicity test: Four isolates of F. oxysporum were selected
were selfed seeds of five cucumber (Cucumis sativus L) (depending on differences in their culture’s colors which varies
inbred lines. These inbred lines were chosen based on their from white to pink with a purple tinge (Federation of British Plant
morphological characters and fusarium wilt disease (Fusarium Pathologists, 1973) to be tested for their pathogenic abilities. In
oxysporum f. sp. cucumerinum) resistance. The five inbred lines this respect, Koch’s postulates were carried out to confirm the
were kindly supported from the Vegetables Breeding Department, virulence of the tested F. oxysporum on a susceptible cucumber
Horticulture Research Institute, Agriculture Research Center, cultivar (cv. Beta Alpha) under pot experiment at the Plant
Giza, Egypt. These inbred lines were Kaha1(P1), Dokky1 (P2), Pathology Research Institute,Agricultural Research Center (ARC),
Dokky2 (P3), Kaha2 (P4) and Dokky3 (P5). Giza, Egypt.
Evaluation for yield and its components and Fusarium Seeds of cucumber (cv. Beta Alpha) obtained from Horticulture
resistance under field conditions: This experiment was carried Research Institute, Agriculture Research Center, Giza, Egypt
out at Kaha Experimental Research Farm of Horticulture were surface sterilized by submersion into solutions of 20-30 %
Research Institute, Qalyubia Governorate. This work was done commercial Clorox (5.25 %) sodium hypochlorite for 1 min,
in previously known naturally infested field with fusarium wilt then washed several times in sterilized distilled water and dried
disease. Half diallel cross design for 5 inbred lines to produce 10 between two folds of sterilized filter paper. Seeds were sown in
hybrids (without reciprocal) was done. The experiment was in plastic trays filled with autoclaved peat-moss substrate.
Journal of Applied Horticulture (www.horticultureresearch.net)
 Breeding for fusarium wilt resistance and some economic characters in cucumber 257

Fusarium isolates were multiplied on sand corn medium according to Antioxidant activity- DPPH-free radical scavenging
Leslie and Summerell (2006). Sand and corn grains were mixed (Inhibition %): The free radical scavenging activity (DPPH)
at the rate of 1:3 (25.0 g clean washed sand; 75.0 g corn grain), was determined by the method described by Begum et al. (2018).
moistened to 50 percent moisture content, filled in 500 mL glass
Statistical analysis: The data of all the quantitative traits were
bottles and then autoclaved at 120 ºC for half an hour. Fungal discs
subjected to analysis of variance (ANOVA) as outlined by Gomez
(5 mm) of pure cultures of each of the tested fusarium isolates
were used to inoculate the prepared sand corn grain bottles then and Gomez (1984). Regression analysis was carried out by using
incubated at 25± 2 °C for 15 days. Pathogenicity test was carried (SPSS software, ver. 22.0, Inc., NY, USA). Also, mean values of
out using autoclaved clay loam soil. Pots (30-cm diameter) were parents and crosses were used to estimate heterosis over mid (MP)
sterilized by immersing in 5 % formalin solution for 15 min and and better parent (BP) as described by Matzinger et al. (1962)
then left for 2 days to insure complete formalin evaporation, the and Fonseca and Patterson (1968), respectively. For the traits
pots containing 5 kg of sterilized soil were infested with the with significant genotypic variances, Griffing’s method 2 model 1
previously prepared inoculum at the rate of 3 % (w/w) watered for (parents and F1’s without reciprocal) (Griffing, 1956) was used to
7 days to enhance growth and distribution of the fungal inoculum estimate general combining ability (GCA) for the five parents and
then, after 30 days of sowing, seedlings were transplanted into the specific combining ability (SCA) for their ten hybrids as outlined
infested potted soil. Each pot was planted by three seedlings. The by Singh and Chaudhary (1985).
pots were irrigated two times weekly. Each treatment included
five replicates. The treatments were arranged in a completely
Results and discussion
randomized block design. Plants were observed for cucumber Evaluation of cucumber genotypes for yield and its components
wilt symptoms. After 60 days of inoculation with pathogen the and Fusarium resistance under field conditions experiment:
disease severity was recorded (Manzoor et al., 2019). Mean performance for some economic traits and disease severity
of five parents and their ten F1’s is presented in Table 1. Data
Host range of highest virulent isolate (F. oxysporum No. 3):
showed that the parental lines for plant length ranged from 71.66
This trial was done to determine the formae speciales under
cm (P1 ) to 180.00 cm (P3 ), while plant length of the hybrids
pot experiment. Plant species belonging to three different
ranged from 100.0 cm (P1×P4) to 175.0 cm (P2×P3). Concerning
families were evaluated against F. oxysporum (No. 3). The three
fruit length, inbred lines ranged from 13 cm (P3) to 20.00 cm (P5)
different families were: A-Solanaceae: tomato (cv. Super Strain
and hybrids ranged from 11.67 cm (P3×P5) to 18.97 cm (P1×P2).
B) and pepper (cv. California). B-Cucurbitaceae: squash (cv.
Early yield ranged from 526.2 g (P1) to1456.7 g (P2), meanwhile,
Eskandrane), watermelon (cv. Sugarbaby) and cucumber (cv.
hybrids ranged from 594.33 g (P2×P3) to 3986.3 g (P1×P3). Parental
Beta Alpha). C-Cruciferaceae: cauliflower (cv. White Magic)
lines ranged from 2911.7 g (P1) to 8348.3 g (P2), while, the hybrids
and cabbage (cv. Balady). Seeds of cauliflower (cv. White Magic)
ranged from 4020.3 g (P1×P2) to 10456 g (P1×P3) for total yield.
were obtained from Sakata Seeds Company, while the rest of
Results in Table 1 show that disease severity for genotypes could
all seeds were obtained from Horticulture Research Institute,
be divided into highly resistant inoculated genotypes, viz.,
ARC, Egypt. F. oxysporum inocula used for inoculation and the
P1×P3, P1×P4, P4×P5, P1×P5, P2 and P2×P4. Three genotypes, viz.,
transplants of different families were prepared as mentioned
P2×P5, P2×P3 and P3×P5 were rated as resistant for this disease.
before. The reaction of the tested plants was indicated by (+) for
While, P4, P3 and P3×P4 were categorized as moderately susceptible
the infected host plants and (-) for the non-infected ones.
genotypes. Genotypes P5 and P1×P2 were recorded as susceptible and
Evaluation of cucumber genotypes for Fusarium wilt resistance only one genotype (P1) was highly susceptible.
under pots experiment: F. oxysporum f.sp cucumerinum (isolate
Correlation among field parameters, economic characters and
No.3) was exploited for evaluating the resistance or susceptibility
disease severity under field conditions: Data in Table 2 revealed
of cucumber genotypes to fusarium wilt. The test was done in pot
that disease severity under field condition was negatively correlated
experiment at the Plant Pathology Research Institute, Agricultural
with early yield (r = -0.619, P ≤ 0.05) and total yield (r =-0.988,
Research Center (ARC), Giza. The inoculum was prepared as
P ≤ 0.01) indicating that total yield is more sensitive to diseases
mentioned before in pathogenicity test. Seeds of fifteen cucumber
severity. However, disease severity was not correlated with any of
genotypes prepared as mentioned before in pathogenicity test.
the other traits. It is worth mentioning that total yield and early
Assessment of cucumber genotypes for different defense yield were positively correlated (r = 0.650, P ≤ 0.01), which
responding biochemical compounds under pot experiment: indicates that early yield could be used to predict total yield trait.
Lower stem of cucumber genotypes were taken when cucumber
Heterosis: Estimates of heterosis over both mid and better
wilt symptoms were observed.
parents for some economic characters in 10 cucumber hybrids
Total phenol content (TPC): Total phenols were measured using is illustrated in Table 3. For early yield and total yield traits,
spectrophotometer at 520 nm against a reagent blank according heterosis values over mid and better parents were highly positive
to Singleton et al. (1999). significant in all studied hybrids except P2×P3 and P3×P4 for early
and total yield, respectively, and P1×P2 for both traits. These results
Polyphenol oxidase (PPO) and peroxidase (POX) activity:
indicated that using these materials are suitable for hybridization
Polyphenol oxidase and peroxidase activity were determined
in developing high yielding hybrids of cucumber that can adapt
according to Matta and Dimond (1963) as absorbance at 420 and
to the zone. These results are in consonance with the findings
425 nm/min/g fresh weight, respectively by spectrophotometer .
of Munshi and Verma (1997) in muskmelon, Chaubey and Ram
Catalase (CAT) activity: CAT activity was determined using the (2004) in bitter gourd and Sarkar (2003) and Munshi et al. (2005)
method described by Aebi (1984). in cucumber. Also, highly negative significant heterosis in most
Journal of Applied Horticulture (www.horticultureresearch.net)
258 Breeding for fusarium wilt resistance and some economic characters in cucumber

Table 1. Mean performance for some economic traits and disease severity of five parents and their ten F1’s in cucumber
Genotype Plant length Branch Internode Internode
Fruit length Fruit diameter Fruit weight Early yield Total yield (D.S.)
(cm) number length (cm) number (cm) (cm) (g) (g) (g) (%)
Parents
P1 71.667 d 146.67 d-f 8.33 ab 6.67 b-e 14.00 c-e 3.00 ab 28.67 bc 526.2 k 2911.7 o 45.06 gh
P2 102.50 cd 190.67 b-d 6.67 ab 9.07 a-c 17.00 a-d 3.25 ab 28.67 bc 1456.7 fg 8348.3 e 11.55 ab
P3 180.00 a 221.67 a-c 8.00 ab 9.50 ab 13.00 de 4.00 a 28.67 bc 1100.0 g-i 5742.3 k 27.10 e
P4 105.00 cd 235.33 ab 8.33 ab 10.67 a 17.33 a-c 2.80 ab 29.67 a-c 1008.5 h-j 6023.0 j 25.12 ef
P5 150.00 a-c 193.33 b-d 5.00 ab 10.17 ab 20.00 a 3.50 ab 29.33 bc 665.00 jk 5139.0 m 32.18 lm
Hybrids
P1×P2 109.33 cd 183.33 b-e 3.67 ab 7.17 a-e 18.97 ab 2.73 ab 31.00 ab 809.67 i-k 4020.3 n 37.45 l
P1×P3 115.33 b-d 181.33 b-e 9.67 a 6.67 b-e 15.58 b-e 2.70 ab 31.67 ab 3986.3 a 10456 a 7.12 ab
P1×P4 100.00 cd 156.67 c-f 4.67 ab 5.33 d-f 14.23 c-e 2.25 b 30.33 ab 2486.3 cd 9877.3 b 7.93 a-c
P1×P5 127.67 a-d 140.67 d-f 3.67 ab 4.00 ef 16.40 a-d 3.47 ab 27.67 bc 3205.7 b 8962.3 d 10.08 ab
P2×P3 175.00 ab 99.00 f 3.67 ab 7.17 a-e 13.00 de 3.75 a 31.00 ab 594.33 k 7831.7 h 18.12 d-f
P2×P4 153.33 a-c 118.67 ef 3.33 b 4.83 ef 15.26 b-e 3.60 ab 21.00 c 1842.0 e 8332.0 f 14.25 b-d
P2×P5 130.00 a-d 279.33 a 4.00 ab 8.50 a-d 17.17 a-c 2.82 ab 38.67 a 2233.7 d 7997.0 g 16.45 c-f
P3×P4 110.67 cd 161.67 c-f 4.33 ab 5.83 c-f 15.17 b-e 3.37 ab 28.33 bc 2706.3 c 5363.0 l 28.15 ef
P3×P5 125.00 a-d 97.00 f 6.33 ab 3.00 f 11.67 e 4.00 a 23.33 bc 1582.3 ef 7241.7 i 19.11 d
P4×P5 152.33 a-c 118.00 ef 2.67 b 3.67 ef 17.77 a-c 2.70 ab 27.33 bc 1196.0 gh 9034.0 c 8.99 a-c
Means within a column followed by the same letter are not significantly different (P = 0.05) according to Duncan’s multiple range test. P1 = Kaha1,
P2 = Dokky1, P3 = Dokky2, P4 = Kaha2, P5 = Dokky3, (D.S.) % = Disease severity Average two seasons
Table 2. Correlation among field parameters economic traits in cucumber under field condition
Correlation   Early yield/ Total yield/ Fruit Fruit Internode Internode Branch Plant Fruit D.S.
m2 m2 diameter length number length number length weight (%)
Early yield\ m2 1.00 0.65**
(a)
-0.23
(b)
-0.09 0.10 -0.37 0.07 0.05 -0.18 -0.62*
Total yield\ m 2
1.00 -0.20 -0.11 0.01 -0.37 -0.16 -0.17 0.21 -0.99**
Fruit diameter 1.00 -0.44 -0.48 -0.03 -0.01 -0.30 0.57*
(c)
0.15
Fruit length 1.00 0.30 0.38 -0.26 0.44 -0.10 0.09
Internode number 1.00 0.49 0.07 0.69 **
-0.13 0.04
Internode length 1.00 0.42 0.77** 0.03 0.37
Branch number 1.00 0.32 -0.35 0.20
Plant length 1.00 -0.13 0.17
Fruit weight 1.00 -0.23
(D.S.) % 1.00
Liner correlation Coefficient (r) is significant at P ≤ 0.01 (**), (b) Liner correlation Coefficient (r) is significant?, (c) Liner correlation Coefficient (r)
is significant at P ≤ 0.05 (*), (D.S.) % = Disease severity in field
of the studied hybrids was observed for plant length, branch genotypes P1 and P3 were good combiners for early yield traits,
number and internode length traits over mid and better parents. meanwhile, the genotypes P2 , P4 and P5 were good combiners for
These negative sign is desired for plant breeder. Three (P1×P3, total yield traits.
P1×P4 and P1×P5) out of ten hybrids showed the highest values of
Specific combining ability (SCA) effect on the vegetative growth,
heterosis over better parent for early yield and total yield traits,
fruit quality and yield and yield components traits of studied
meanwhile, the hybrid P1×P3 was the best one in previous traits plus
ten hybrids are presented in Table 5. SCA reveals the best cross
fruit length trait. Similar trend was observed for fruit weight and
combinations which can be useful for developing hybrids with
total yield among the same three hybrids gave the highest values
high vigor for the traits. Significant superior SCA effects for all
for both mid and better parents, respectively. Singh et al. (2013)
studied traits were not shown by a single hybrid. Data obtained
found high positive heterosis for seed yield/ plant in bitter gourd.
in Table 5 indicated that the F1 crosses P1×P3 and P2×P5 achieved
General and specific combining ability: Combining ability is significant positive SCA effects for plant length. Only one hybrid
useful in successful prediction of genetic capability of parental (P1×P3) showed significant SCA effect for branch number. Also,
lines and crosses. General combining ability (GCA) of cucumber only one cross (P2×P5) showed significant SCA effect for internode
growth and fruit yield traits showed that genotype P1 had positive length. Two crosses (P1×P3 and P2×P5) showed significant SCA
GCA effect for branch number and early yield traits (Table 4). effects for internode number. Three hybrids exhibited significant
Furthermore, it was observed that GCA of P2 was significantly SCA effects for fruit length and the cross P1×P2 showed the highest
positive for plant length, internode length, fruit length and total significant value. The hybrids P1×P5 and P2×P4 showed significant
yield traits. Genotype P3 had significantly positive GCAfor branch SCA effects for fruit diameter. The hybrid P2×P4 showed the highest
number, fruit diameter, fruit weight and total yield characters. significant SCA effect for fruit weight. Six crosses (P1×P3, P1×P4,
For genotype P4, significantly positive GCAwas observed for only P1×P5, P2×P4, P2×P5 and P3×P4) had significant positive SCA effects
total yield trait. Genotype P5 showed significantly positive GCA for early yield. For total yield, the SCA effects for eight hybrids
for fruit length, fruit weight and total yield traits. In general, the (P1×P3, P1×P4, P1×P5, P2×P3, P2×P4, P2×P5, P3×P5 and P4×P5) were
Journal of Applied Horticulture (www.horticultureresearch.net)
 Breeding for fusarium wilt resistance and some economic characters in cucumber 259

Table 3. Heterosis values over mid (MP) and better (BP) parents for some economic characters in 10 cucumber hybrids
Hybrids Plant Branch Internode Internode Fruit Fruit Fruit Early Total
length number length number length diameter weight yield yield
Heterosis over mid parents (%)
P1×P2 8.69 -51.10** -8.90 8.14 22.30** -12.53 25.50 -18.30** -28.50**
P1×P3 -1.54** 18.30* -17.53** 10.46 15.46 -22.86 -8.35 390.20** 141.60**
P1×P4 -17.97 -43.90** -38.40** 3.99 -9.15 -22.40** 13.20** 224.00** 121.20**
P1×P5 -17.25** -44.90** -52.48** -4.59 -3.53 6.67 15.19 438.20** 122.60**
P2×P3 -51.90** -49.90** -22.80** 8.14 -13.30** 3.45 23.80** -53.50** 11.10**
P2×P4 -44.20** -55.50** -51.01** -28.00** -11.10** 19.01 47.70** 49.44** 15.90**
P2×P5 45.40** -31.43 -11.61* 33.33** -7.21 -16.54 2.97 110.50** 18.50**
P3×P4 -29.20** -46.90** -42.15** -2.85 0.003 -0.98 -22.30** 156.70** -8.83**
P3×P5 -53.20** -2.56 -69.40** -19.54** -29.20** 6.67** -24.24 79.30** 33.10**
P4×P5 -44.90** -59.90** -64.80** -7.35 -4.80** -14.20** 19.47 42.93 61.80**
Heterosis over better parents (%)
P1×P2 -3.85 -55.90** -20.96** 8.14 11.50** -15.90 6.66 -44.70** -51.80**
P1×P3 -18.10** 16.00 -29.82** 10.46 11.30** -32.50** -35.90** 262.30** 82.10**
P1×P4 -33.40** -44.00** -50.00** 2.25 -17.80** -25.00 -4.76* 146.50** 63.90**
P1×P5 -27.20** -55.99 -60.60** -5.68 -18.00** -0.95 -14.89 382.10** 74.40**
P2×P3 -55.30** -54.00** -24.50** 8.14 -23.53 -6.25 -2.78 -59.40** -6.19**
P2×P4 -49.50** -60.00** -54.69** -29.21** -11.90** 10.77** 46.00** 25.70** -0.20**
P2×P5 44.40** -40.00 -16.40** 31.8** -14.10** -19.50 -13.33 52.40** -4.20**
P3×P4 -31.30** -48.00** -45.31** -4.50 -12.50** -92.35 -38.50** 146.00** -10.90**
P3×P5 -56.20** -20.83 -70.49** -20.45** -41.67 0.00 -30.50** 43.80** 26.10**
P4×P5 -49.80** -67.90** -65.63** -7.87 -11.17 -22.86 1.55** 18.50** 49.90**
* and ** significant at 5 % and 1 % levels of probability, respectively. P1 = Kaha1, P2 = Dokky1, P3 = Dokky2, P4 = Kaha2 and P5 = Dokky3.
Table 4. General combining ability (GCA) effects for some economic characters in five parental genotypes of cucumber
Parents Plant Branch Internode Internode Fruit Fruit Fruit Early Total
length number length number length diameter weight yield yield
P1 -7.71** 0.77** -0.63** 0.54 -0.20 -0.29** -23.92** 197.25** -538.3**
P2 7.48** -0.70** 0.70** 0.70 0.54** 0.03 1.36 -252.37** 280.83**
P3 -3.86 1.01** 0.11 -0.35 -1.89** 0.38** 17.55** 129.96** -76.41**
P4 2.33 -0.18 0.02 -1.11** 0.35 -0.24** -5.26 12.58 248.63**
P5 1.76 -0.90** -0.20 0.22 1.20** 0.12 10.27** -87.42** 85.88**
S.E. (ĝi- ĝj) 7.31 0.60 0.35 0.89 0.43 0.15 6.21 37.23 0.76
* and ** Significant at 5 % and 1 % levels of probability. P1 = Kaha1, P2 = Dokky1, P3 = Dokky2, P4 = Kaha2 and P5 = Dokky3
significant and positive. Among all hybrids, P1×P3, P1×P4, P1×P5, who observed that better performing crosses had at least one of the
P2×P4 and P2×P5 exhibited significant SCA effects for both early cultivars with high GCA effect. According to Singh et al. (2013),
and total yield characters. So, these hybrids can be used in future high × low general combining ability combinations are suitable
breeding program to improve these traits. for heterosis breeding, while, high × high general combining
ability combinations can be considered for developing superior
The crosses with high significant specific combining ability
variants through pedigree method. The estimates of GCA effects
effects are useful to obtain high performing hybrids. This could
(Table 4) revealed that none of the parents exhibited good GCA
be because they involved parents with high × high, high × low and
for all studied characters. So, it was difficult to pick up good
low × low general combining ability effects which indicated the
combiners for all traits together because the combining ability
presence of additive and dominance gene effects for controlling
effects were not consistent for all yield components (Solanki
these traits. Also, the superiority of cross combinations involving
and Shah, 1990). This shows that genes for different desirable
high × low or low × low general combiners as parents may be characters would have to be combined from different sources
attributed to the genetic diversity among parents, in the form (Nehe et al., 2007).
of number of heterozygous loci of the parents involved in the
cross combinations. Some of the parents with high GCA effects Phenotypic correlation: The phenotypic correlation coefficients
produced hybrids with low SCA effects. This may be due to lack of among nine cucumber traits are presented in Table 6. Highly
complementation of the parental genes. On the other hand, parents positive significant phenotypic correlation was recorded between
with low GCA effects produced hybrids with high SCA effects total yield and early yield. Also, highly positive significant
which can be attributed to complementary gene effect. Similar phenotypic correlation were found among plant length and both
results were reported by Laxuman et al. (2012) and Singh et al. internode number and internode length. Positive significant
(2013) in Momordica charantia L., Hussien and Hamed (2015) phenotypic correlation between fruit weight and fruit diameter
in pumpkin and Reddy et al. (2013) in Abelmoschus esculentus was found. These results are in agreement with the findings of
Phani et al. (2018).
L. Moench. Furthermore, crosses with good specific combining
ability had one of its cultivars as good general combiner of that Cluster Analysis: Genetic divergence studies in cucumber
trait. This was similar to the findings of Choudhary et al. (2000) revealed some interesting features of differentiation and
Journal of Applied Horticulture (www.horticultureresearch.net)
260 Breeding for fusarium wilt resistance and some economic characters in cucumber

adaptability such as cluster analysis which provide additional basis by implementing crossing. The cluster analysis obtained
information for studying interrelationship between genotypes from UPGMA classified the fifteen (five parents and their 10 F1
and giving graphical assessment of genetic variability. For this hybrids) cucumber genotypes to five clusters (Fig. 1). The first
purpose hierarchical cluster analysis, on the basis of average group (A): consists of four hybrids, viz., P2×P4 (11), P3×P5 (14),
linkage (within groups) and interval Euclidean distance, was
P1×P4 (8) and P2×P5 (12). The second group (B): contains five
applied to investigate genetic distance and diversity between
parents plus two hybrids, viz., P3 (3), P1×P2 (6), P1 (1), P2 (2), P4 (4),
the five parents and their 10 F1 hybrids. The data matrix of the
dissimilarity coefficient on the basis of Euclidean distance is P5 (5) and P4×P5 (15). While, both the third (C) and fifth groups
presented in Table 7. Wide range of genetic distance among (E) have a unique hybrid for each one P2×P3 (10) and P3×P4 (13),
studied genotypes reflected the presence of wide range of genetic respectively. The fourth group (D) had two hybrids, viz., P1×P3
variation and provides an opportunity to improve the genetic (7) and P1×P5.
Table 5. Specific combining ability (SCA) effects for some economic characters in ten hybrids of cucumber
Hybrids Plant Branch Internode Internode Fruit Fruit Fruit Early Total
length number length number length diameter weight yield yield
P1×P2 15.35 -1.89** 0.28 0.73 2.86** -0.21 4.71 -828.47** -2873.5**
P1×P3 24.68** 2.40** 0.37 2.44** 1.91** -0.58** -5.48 1965.86** 3919.37**
P1×P4 -6.17 -1.41 -0.87** 1.87 -1.6** -0.42** 1.99 583.24** 3015.65**
P1×P5 -21.60** -1.70** -1.99** -2.13* -0.36 0.44** 14.13* 1402.57** 2263.41**
P2×P3 -72.84** -2.13** -0.46 1.63 -1.4** 0.14 28.90** -976.52** 475.27**
P2×P4 -59.37** -1.27 -2.70** -7.60** -1.4** 0.61** 30.04** 388.52** 650.56**
P2×P5 101.87** 0.11 1.18** 8.73** -0.34 -0.53** -8.82 880.19** 478.32**
P3×P4 -5.03 -1.98** -1.11** 0.78 0.93* 0.03 -28.8** 870.52** -1961.2**
P3×P5 -69.13** 0.73 -3.73** -5.56** -3.4** 0.31 -30.0** -153.48** 80.22**
P4×P5 -54.32** -1.75** -2.97** -0.80 0.45 -0.37** 20.14** -422.43** 1547.51**
SE(Sij-Sik) 17.91 1.47 0.85 2.19 1.05 0.36 15.22 91.21 1.86
SE(Sij-Skl) 16.35 1.34 0.78 2.00 0.96 0.32 13.89 83.26 1.69
* and ** Significant at 5 % and 1 % levels of probability. P1 = Kaha1, P2 = Dokky1, P3 = Dokky2, P4 = Kaha2 and P5 = Dokky3
Table 6. Phenotypic correlation coefficients among some economic characters in cucumber
Traits   Early Total Fruit Fruit No Internode Branch Plant
yield yield diameter length node length number length
Total yield 0.650**              
Fruit diameter -0.232 -0.203            
Fruit length -0.086 -0.110 -0.438          
No node 0.104 0.012 -0.482 0.304        
Internode length -0.370 -0.372 -0.032 0.378 0.493      
Branch number 0.071 -0.155 -0.009 -0.261 0.069 0.419    
Plant length 0.050 -0.170 -0.299 0.438 0.690** 0.766** 0.324  
Fruit weight -0.178 0.213 0.572* -0.104 -0.127 0.035 -0.349 -0.132
*and ** Significant at 5 % and 1 % levels of probability.
Table 7. Genetic distance among five cucumber genotypes and their ten F1 hybrids based on some economic characters
Genotypes Rescaled Squared Euclidean Distance
P1 P2 P3 P4 P5 P1×P2 P1×P3 P1×P4 P1×P5 P2×P3 P2×P4 P2×P5 P3×P4 P3×P5 P4×P5
1 P1 1.00
2 P2 0.01 1.00
3 P3 1.00 1.00 1.00
4 P4 1.00 1.00 1.00 1.00
5 P5 0.02 0.01 0.02 0.01 1.00
6 P1×P2 1.00 0.01 1.00 0.01 0.03 1.00
7 P1×P3 0.23 0.23 0.20 0.25 0.35 0.18 1.00
8 P1×P4 0.04 0.03 0.02 0.04 0.09 0.02 0.09 1.00
9 P1×P5 0.18 0.18 0.15 0.20 0.29 0.14 1.00 0.06 1.00
10 P2×P3 0.07 0.05 0.08 0.05 0.02 0.09 0.51 0.17 0.43 1.00
11 P2×P4 0.02 0.01 0.01 0.02 0.05 0.01 0.14 0.01 0.10 0.11 1.00
12 P2×P5 0.06 0.06 0.04 0.07 0.12 0.03 0.06 0.01 0.04 0.23 0.02 1.00
13 P3×P4 0.57 0.59 0.53 0.62 0.77 0.50 0.08 0.35 0.12 1.00 0.44 0.28 1.00
14 P3×P5 0.02 0.01 0.01 0.02 0.05 0.01 0.14 0.01 0.11 0.11 1.00 0.02 0.45 1.00
15 P4×P5 0.02 0.01 0.02 0.01 1.00 0.03 0.34 0.08 0.28 0.02 0.04 0.12 0.75 0.04 1.00

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 Breeding for fusarium wilt resistance and some economic characters in cucumber 261

Morphological identification of the isolated fungi: Four fungal from 20.27-30.50 and 5.00-6.75 μm. Clamydospores were present
isolates were selected depending on the differences in their and formed singly or in pairs (Fig. 2). Isolates were identified as
culture’s colors (Federation of British Plant Pathologists, 1973). F. oxysporum due to occurrence of typical macroconidia with
These isolates were identified as F. oxysporum (named 1, 2, 3 and foot-shaped basal cells, microconidia borne in false heads only
4). Microscopic observations revealed that the mycelia of the on monophialides with oval shaped and chlamydospores (Owen,
isolates were delicate, white to pink with a purple tinge, sparse to 1956 and Martínez et al., 2003).
abundant than floccose, margins slightly lobed or smooth on
PDA. Microconidia formed singly, oval to reniform and without Pathogenicity test: Four isolates of F. oxysporum were tested for
any septation. The size of microconidia ranged from 7.50-10.25 their pathogenic activities on cucumber susceptible cultivar (cv.
and 2.50-3.50 μm. Conidiogenous cells bearing micro- and Beta alpha) as shown in Table 8. In this respect, the F. oxysporum
macroconidia were monophialides type. Macroconidia were isolate No.3 gave the highest disease severity (52.38 %) followed
falcate to almost straight, the size of the macroconidia ranged by isolate No.4 (47.78 %) without significant differences between
them. On the other hand, isolate No.1 (29.58 %) and isolate No.2
(32.41 %) recorded the lowest ones without significant differences
between them.
A Table 8. Pathogenicity test of the four isolates of F. oxysporum against
the susceptible cucumber cv. Beta Alpha under pots experiments
Isolates code F. oxysporum Control LSD
1 2 3 4 (P=0.05)
Wilt disease
severity % 29.58 32.41 52.38 47.78 00.0 6.76
B

Due to infection with F. oxysporum, the host plant fails to absorb

C
water and nutrition from the soil, and the resulting process is
called “occlusion of vessels” through the formation of callose,
tylose or gels. Finally, the stomata in the leaves will close and
D wilted, followed by death of the entire plant (Yadeta and Thomma,
2013).
E
Host range: F. oxysporum includes a lot of non-pathogenic
Fig. 1. Dendrogram of five parental and their 10 F1 hybrids based on some strain in addition to host specific pathogenic strains (formae
economic traits. Numbers used for five cucumber genotypes and their ten speciales). Isolate No.3 was selected in this study due to its high
F1 hybrids are: 1:P 1; 2:P 2; 3:P 3; 4:P 4; 5:P 5; 6:P1×P2; 7:P1×P3; 8:P1×P4;
9:P1×P5; 10:P2×P3; 11:P2×P4; 12:P2×P5; 13:P3×P4; 14:P3×P5; 15:P4×P5; pathogenicity in previous trials. Data in Table 9 indicate that no
symptoms were observed on the cruciferaceae (cabbage and
cauliflower), solanaceae (tomato and
pepper) and cucurbitaceae (Squash and
watermelon) hosts. However, only a
typical symptom of fusarium wilt was
detected on cucumber cv. Beta Alpha.
Depending on the results of host range
experiment, it is clear that the main
causal organism of wilting symptoms
observed on cucumber plants is F.
oxysporum f. sp. cucumerinum. Most
species belong to the F. oxysporum
complex with n u m e r o u s special
forms attacking different plant species.
Individual special form susually have a
very restricted host range (Armstrong
and Armstrong, 1981; Ruiz-Roldán and
Di Pietro, 2012) .
Enzymatic activities, total phenol
c o n t e n t s , D P P H - f re e r a d i c a l
scavenging activity and disease
severity in pot experiment: The
v a l u e s o f P P O , P O X a n d CAT
activities differed between genotypes
Fig. 2. Morphological identification of Fusarium oxysporum. A: Colony morphology of Fusarium
oxysporum ( isolate No. 3) on potato dextrose agar (PDA; 14 days old) showing fluffy growth white to as represented in Fig 3. The most
pink aerial mycelia. B: Chlamydospores were present and formed singly or in pairs. C: Macroconidia susceptible genotypes, viz., P1, P3, P5
with foot-shaped basal cells. D: Microconidia oval -shaped , P1×P2 and P3×P4 had the lowest PPO,
Journal of Applied Horticulture (www.horticultureresearch.net)
262 Breeding for fusarium wilt resistance and some economic characters in cucumber

Table 9. Host range of F. oxysporum isolate No.3 under pots experiments Table 10. Effect of infested soil with F. oxysporum f. sp. cucumerinum
on total phenol content (TPC), radical scavenging activity (DPPH) and
Host Cultivar Reaction disease severity (D.S. %) of cucumber genotypes
Tomato Super Strain B - Genotypes TPC (mg/g fresh DPPH D.S.
weight) (IC50mg/mL) (%)
Pepper California wonder -
P1 70.13 101.0 68.16
Squash Eskandrane -
P2 127.6 63.1 14.12
Cucumber Beta Alfa + P3 85.59 95.53 33.3
Watermelon Sugarbaby - P4 87.49 88.87 32.43
Cauliflower White Magic - P5 83.31 100.05 45.23
Cabbage Balady - P 1 × P2 71.32 102.21 55.2
P 1 × P3 135.45 51.1 9.15
POX and CAT activities associated with highest disease severity P 1 × P4 133.58 54.2 10.23
(%). On other hand, the most resistant genotypes P2, P1×P3, P1×P4, P 1 × P5 129.63 61.7 13.43
P1×P5 and P4×P5 had the highest PPO, POX and CAT activities 79.6
P 2 × P3 93.13 24.7
associated with lowest disease severity (%).The results indicated 67.2
P 2 × P4 116.43 20.21
that there were remarkable increase in the enzyme activities in
P 2 × P5 100.0 78.43 22.72
cucumber plants depending on different models of genetic disease
P 3 × P4 84.85 96.15 35.4
resistance. This is in conformity with the finding of Khatun et
P 3 × P5 92.32 84.3 25
al. (2009) and Huang et al. (2019). Peroxidase and polyphenol
P 4 × P5 130.79 60.1 12.88
oxidase are involved in the oxidation of specific host metabolites
which in turn act as inhibitors of growth of the phytopathogen L.S.D ( P≤0.05) 0.74 2.01 3.45
(Begum et al., 2018). P1 = Kaha1, P2 = Dokky1, P3 = Dokky2, P4 = Kaha2 and P5 = Dokky3
recorded 85.59, 84.85, 83.31, 71.32 and 70.13 %, respectively.
The antioxidant enzyme CAT showed contrary activities in
resistant and susceptible cultivars. Both CAT and peroxidase are Total phenol content (TPC) plays an important role in the
antioxidant enzymes eliminating reactive oxygen species (ROSs). mechanism of disease resistance. They accumulated rapidly
Catalase catalyzes elimination of H2O2 during oxidative damages during host pathogen interactions and mediated disease
(Luhova et al., 2006). Thus, the patterns and levels of the activity suppression through participating in the oxidation of TPC to
of both enzymes in the resistant cultivar may be correlated with quinines, leading to the increase in antimicrobial activity and
defense responses related to H2O2 accumulation. Additionally, inhibiting pathogen progression. Then disease severity was
H2O2 induces cell death and acts as a signal for induction of
negative and highly correlated with the biochemical parameters
systemic defense responses (Heller and Tudzynski, 2011).
particularly in phenolic compounds, which can be used to predict
As can be seen from the results presented in Table 10, infection disease severity as shown in Fig. 4. This could reduce the amount
of cucumber plants with F. oxysporum isolate No.3 caused of work required in screening tests of disease resistance (Khatun
significant decrease in disease severity and significant increases et al., 2009).
in total phenol contents in case of genotypes P1×P3, P1×P4, P4×P5,
P1×P5 and P2, the percentages of increase in the contents of Scavenging activities of cucumber genotypes varied from 51.1
total phenol were 135.45 , 133.58 , 130.79 , 129.63 and 127.60 to 101.0. Genotype P 1×P 3 gave the highest content of total
%, respectively. whereas, the lines P3, P3×P4, P5, P1×P2 and P1 phenolic contents (TPC) (135.45mg/100 g), which causes its

Fig. 3. Effect of infested soil with Fusarium oxysporum f. sp. cucumerinum on catalase (CAT), peroxidase (POX), poly phenol oxidase (PPO) activities
and disease severity (D.S. %) of cucumber genotypes.
Journal of Applied Horticulture (www.horticultureresearch.net)
 Breeding for fusarium wilt resistance and some economic characters in cucumber 263

Fig. 4. Regression equation that describes the effect of total phenol on

disease severity in cucumber genotypes
stronger antioxidant ability as DPPH radical scavenging activity
(IC50 51.1 mg/mL) followed by P1×P4 (IC5054.2 mg/mL) and
P4×P5 (IC5060.1 mg/mL), whereas P1 caused weak antioxidant
ability as DPPH radical scavenging activity (IC50101.0 mg/mL). Fig. 5. Regression equation that describes the effect of total phenol on
Furthermore, there was a high correlation between the TPC and radical scavenging activities (DPPH).
the scavenging activity. Therefore, the presence of phenolic Chaubey, A.K. and H.H. Ram, 2004. Heterosis for fruit yield and its
compounds in different genotypes contributes significantly components in bitter gourd (Momordica charantia L.). Veg. Sci.,
to their antioxidant potential as shown in Fig. 5. Antioxidant 31(1): 51-53.
properties of phenolic compounds are directly linked to their Choudhary, A.K., L.B. Chaudhary and K.C. Sharma, 2000. Combining
structure. Indeed, phenolics are composed of one (or more) ability estimate of early generation inbred lines derived from two
maize populations. Ind. J. Genet., 60(1): 55-61.
aromatic rings bearing one or more hydroxyl groups and are
Difco manual of dehydrated culture media and reagents for microbiological
therefore potentially able to quench free radicals by forming and clinical laboratory procedures. Detroit1, Michigan, 9 ed.1985.
resonance-stabilized phenoxyl radicals (Rice-Evans et al., pp 364.
1996; Bors and Michel, 2002). Ene, C.O., P.E. Ogbonna, C.U. Agbo and U.P. Chukwudi, 2019. Heterosis
and combining ability in cucumber (Cucumis sativus L.). Information
From this study, it can be concluded that the lines P2, P4 and P5 Processing Agriculture, 6: 150-157.
showed significant positive GCA effects for total yield trait under Federation of British Plant Pathologists, 1973. A guide to the use of terms
fusarium wilt stress. So, these parents could be successfully used in plant pathology. Phytopathology, pp. No. 17.
in future breeding programs. Also, among all crosses, P1×P3, P1×P4, Fonseca, A. and F.L. Patterson, 1968. Hybrid vigour in a seven-parent
P1×P5, P2×P4 and P2×P5 exhibited significant SCA effects for both diallel cross in common winter wheat (T. aestivum L.). Crop Sci.,
8: 85-88
early and total yield characters under fusarium wilt stress. So, these
Glover, M.A., D.B. Willmot, L.L. Darrah, B.E. Hibbard and X. Zhu,
hybrids can be used in future breeding program.
2005. Diallel analyses of agronomic traits using Chinese and US
maize germplasm. Crop Sci., 45(3):1096-1102.
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